Living at the edge: local versus positional factors in the long-term population dynamics of an endangered orchid.

Populations at the margin of geographic ranges of distribution have been considered more vulnerable than central ones, but recent reviews have caste doubt on this generalization. We examined the reproductive and demographic performance of a rare Euroasiatic orchid (Cypripedium calceolus) at its southwesterly range limit and compared our findings with those of previous studies of nine central populations at the center of the orchid's range. We sought to test the central-marginal model and to evaluate factors involved in long-term performance of forest Eurosiberian species with peripheral populations in southern European mountains. We characterized (structure, temporal fluctuations, herbivory, reproductive success, and recruitment at different habitats) four Pyrenean populations of C. calceolus of different sizes (5-3500 ramets) and monitored three of them for up to 13 years. Two quantitative stochastic models (count data and matrix models) were used to assess population trends and viability and the effect of herbivory. Contrary to expectations, and despite the negative effect of sporadic events of herbivory, the peripheral populations we studied (except the smallest one) performed similarly or better than populations occurring in central part of the species' range in terms of reproductive success and population growth rates. Landscape changes over the last 50 years suggest that natural reforestation could be involved in the success of this plant at its southern limit. Forest expansion in the mountain regions of southern Europe may provide new opportunities for plants with geographic distributions centered mainly at higher latitudes and give some hope for their recovery in future scenarios dominated by biodiversity loss.